Tree vascular tissues connect resource availability to tree physiological processes and growth. The xylem transports water from the soil up to the canopy of even 100-metre tall trees, whereas phloem transport connects the photosynthesis in leaves and the tree metabolic processes, including growth and tree defences against insect and pathogen attacks. Water deficit results in the closing of leaf stomata and decreasing photosynthetic production, as water and carbon dioxide are exchanged through the stomata between the leaf and ambient air. Phloem transport is driven by turgor pressure gradients generated by the interplay of phloem osmotic concentration and xylem water potential. Trees have adapted to local environmental conditions and they adjust to fast environmental changes with physiological responses. This thesis investigates tree physiological responses in vascular tissues, such as osmolality, water potential and stomatal conductance, to environmental conditions in two conifers: Scots pine and Norway spruce.
Seasonality in soil temperature and soil water content affect soil-to-leaf hydraulic conductance, and stomatal conductance is connected to these seasonal patterns in water transport. Soil environment is thus mediated to tree functionality through tree water transport. This thesis also supports Münch’s theory that it is plausible to explain phloem transport in conifers in field conditions with osmotic gradients and gravity. Xylem water potential reflects to osmotic potential and turgor pressure of the inner bark by modifying tissue solute and water content. The turgor gradients hence seem to determine daily and seasonal carbon allocation patterns according to water availability. Pathogenic infections may introduce more rapid changes in tree hydraulic conductance through a decrease in xylem sap surface tension and xylem conductivity during massive invasions of bark beetles that vector blue-stain fungi such as Endoconidiophora polonica. These pest attacks weaken tree vitality and may also increase tree vulnerability to hydraulic failure in the xylem.
A mature tree stem generally consists of a column of wood that is composed of a series of annual incremental layers and enclosed in a covering of bark. The dynamic variations of the bark are complex due to its structure and function: the thick outer-bark acts as a protective barrier against the abiotic and biotic environment; while the phloem is where sugar transport occurs. Much of the bark variation is due to the transport of sugars and its related processes. The xylem pathway, which transports water in the opposite direction, is connected to the phloem in parallel along the entire length of the stem. The immediate connection between these two transport pathways suggests a functional linkage.
The purpose of this thesis is to study the dynamic processes that occur within the bark and its interaction with other internal tree processes and the external environment. These interactions have not been thoroughly quantified, especially on an intra-annual (e.g. daily) scale.
The thesis consists of four papers, of which one is a modelling paper and three are experimental studies. Growth is estimated with the model by separating the water-related influences from measured inner-bark, revealing a growth signal – proxy for cambial stem growth. Using this signal, a correlation study to microclimate variables is examined in one paper; and to assumed growth respiration in a second paper. The remaining two papers explore the seasonality of photosynthesis and respiration, and bark stem dynamics during the spring recovery period.
As a conclusion of this thesis, these papers show how inextricably linked individual tree processes and the environmental are to the changes within the bark. The culmination of this thesis opens new opportunities to further understand the dynamics of bark hydraulics and ecophysiological processes by implementing field measurements and state-of-the-art modelling.